Tuesday, March 19, 2019

Pogonophora - Central position in evolution


Pogonophora: Their spectacular role in animal evolution

A spectacular connection of the two main higher animal groups

Early embryology of the two main lines of animals

     The Pogonophora show how the radial and indeterminate cleavage of deuterostomes (echinoderms, hemichordates and vertebrates) came about from pre-Cambrian annelids in the deep sea while other annelids retained the spiral and determinate cleavage of protostomes as they were giving rise to arthropods and mollusks.

     The deep sea had low input of nutrients that put a premium on not investing in heavy egg shells that protected eggs of protostomes as they were confined in the spiral pattern of cleavage, a confinement that was released by less confining outer membranes that allowed a more direct radial cleavage pattern.  At the same time radial cleavage did not produce the immediate cell fate into a particular tissue, thus making possible more than one viable embryo from a single egg as each of the early cleavage cells retained all of the development potential of the original egg.  The ability to complete development without all the original cleavage products had high survival value in the rigorous deep-sea environment, although details are speculative.

    My comparative study of Tasmanian and Michigan isopods included observation of the impact of the egg membrane’s role in confining development, leading to a unique egg appendage, a clear indication that evolution can occur in developmental stages independent of adult development.  (see more on “Origin of deuterostome embryology” in blog post dated 6/24/2013)

Systems inversion

A simple process

The inverted position of blood vessels, nerves, oral openings of deuterostomes as compared to protostomes (especially the ancestral annelids of both groups) was proposed as evidence of the central role of annelids in evolution of higher animal phyla.  The original reason for rejecting the theory was the drastic difference in early embryology of the two lines of animals.  The next reason thought to negate the annelid theory as well as the embryological argument against it was the use of nucleotide and other molecular data.  Such data must be realigned taking into consideration the effect of the astronomically slow rate of genetic change in the pogonophorans ancestors.

Inversion, the first step

The inversion of annelids began with certain polychaetes that began living vertically in tubes they secreted, as seen in Sabella and many other shallow water polychaetes.   As those growing in progressively deeper water, with less food, became dependent on absorption of nutrients in pore water of the sediments they outcompeted those wasting energy on producing a mouth and some bilateral structure.  This stage is still found in abyssal pogonophorans. 

Inversion, the second step

The return of descendants to shallow seas occurred once the worst episode of pre-Cambrian asteroid bombardment eased.  As they arrived in shallower more nutrient rich areas, they reformed the remnants of their digestive system with a new mouth on the former dorsal side which became the ventral side, as they groped around the sediment surface near their tube, finding food particles that pushed the epidermal and gut layer together triggering mouth formation on the former dorsal surface without the restriction of the nervous system that originally had encircled the esophagus.  Other clues to this step are presented by the parallels of endocrine hormone function, transport, and structural similarities of vertebrates, arthropods, and annelids.

Protostome-Deuterostome links

Segmentation/Metamerism

The segmentation of annelid type was found on a short portion of the most deeply embedded part of some pogonophorans; it included setae that are a very annelid like characteristic.  A few anterior regions are noticeable but without the posterior segmented region it would be difficult to make an annelid connection.  The metamerism of chordates such as ourselves seen in bone, muscle, nervous system and blood vessels is now easily understandable with the intermediate stage of pogonophorans.
The transition from pogonophorans to chordates is best shown by the larval stage comparisons of pogonophorans and hemichordates.

Molecular evidence

Molecular features of several types show greater similarity between deuterostomes and advanced protostomes than their earliest variants found in more ancient protostomes once thought to be the closest common ancestors at the protostome-deuterostome split.

DNA/RNA studies of evolutionary relationships at the phylum level need reevaluation because major ones have ignored the mutation rate differences associated with generation times.  Many well focused studies have shown generation time does affect evolution rates.  One impact has been the Pogonophora showing up in many odd places in phylogenetic trees because they are almost unchanged since their divergence from major groups that have diverged even more from more recent relatives.

Six other posts, from June 17 to June 20, 2013 have additional clarification of the points made above.  The second June 30th post of that year is an annotated bibliography that has some emphasis on protostome/deuterostome comparisons.

Joseph Engemann   
Emeritus Professor of Biology, Western Michigan University, Kalamazoo, Michigan    May 19, 2019

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